Records of the Western Allstralzall Mllsellm Supplement No. 57: 291-298 (1999).

First Australian Pliocene molossid bat: Mormopterus (Micronomus) sp. from the Chinchilla Local Fauna, southeastern Queensland

l 2 Suzanne J. Hand!, Brian S. Mackness , Cecil E. Wilkinson and Doris M. Wilkinson2

1 School of Biological Science, University of New South Wales, NSW 2052; email: [email protected] 29 Birkett Street, Chinchilla, Qld 4413

Abstract - An isolated upper canine from the Pliocene Chinchilla Local Fauna of southeastern Queensland is identified as representing the Australian molossid genus Mormopterus, subgenus Microllomlls. It is the first Australian Pliocene representative of the cosmopolitan family Molossidae and the first Tertiary representative of the Microl1omlls lineage. Approximately six Microl1omlls species are today widely distributed across the Australian mainland, Papua New Guinea and Ambon, and are found in most habitat types from desert to rainforest. The discovery of an indeterminate species of Microl1omlls in the Chinchilla Local Fauna does not contradict the palaeoenvironmental interpretation of the area as woodland savannah.

INTRODUCTION Superfamily Vespertilionoidea Gray, 1821 A variety of taxa has been recovered from the early (Weber, 1928) to middle Pliocene freshwater fluviatile Chinchilla Family Molossidae Gray, 1821 Sand northwest of Chinchilla, southeastern Queensland (Archer 1977, 1982; Archer and Dawson Monnopterns (Micronomus) sp. 1982; Bartholomai 1962, 1963, 1966, 1967, 1971, 1973, 1975, 1976; Bartholomai and Woods 1976; Dawson Material Examined 1982; Flannery and Archer 1983; Gaffney 1981; QM F30575, a left upper canine (Figure 1). Gaffney and Bartholomai 1979; Godthelp 1990; Measurements: Length 1.17 mm, width 0.98 mm, Hutchinson and Mackness submitted; Kemp and height 1.93 mm. Molnar 1981; Mackness and Scanlon in press; Olson 1975, 1977; Patterson and Rich 1987; Woods 1956, Locality and Age 1960, 1962; Wroe and Mackness 1998). Collectively North bank of the Condamine River, this assemblage is known as the Chinchilla Local Chinchilla Rifle Range (26°48'S., 1500 41'E.). The Fauna (Archer and Bartholomai 1978; Woodbume et Chinchilla Sand was named by Woods (1960) for al. 1985; Rich et al. 1991). a sequence of weakly consolidated grey to This paper describes an isolated canine recovered yellowish and light brown sands, ferruginized from the Chinchilla Sand which represents the first heterogeneous conglomerates, grits, sandy clay Australian Pliocene representative of the and clays. These outcrops range from shallow cosmopolitan family Molossidae, and the first beds to sections several metres deep. Tertiary representative of the modern Australian Wilkinson's Quarry has been continuously molossid lineage Micrononws. Two older molossids, worked by two of us (C.E.W., D.M.W.) for over described from Miocene deposits at Riversleigh in ten years. The specimen described comes from a northwestern Queensland, are members of extinct fossil-bearing unit within the quarry that lies molossid lineages (Hand 1990; Hand et aL 1997). unconformably on top of an indurated layer of Taxonomy and dental terminology follow fine sand. The sediments are primarily fluviatile Legendre Hand (1990) and Strahan (1995). in nature and represent a number of depositional The prefix QM 12 indicates specimens registered in events. Most fossils in these units occur as the fossil collection of the Queensland Museum, isolated pieces. Brisbane. Tedford et aL (1992) concluded that the Chinchilla Local Fauna biocorrelates with the Kanunka Local Fauna of the Tirari Desert, South Australia. On the SYSTEMATIC PALAEONTOLOGY basis of magnetostratigraphy, they concluded that the latter is approximately 3.4 million years old, or Suborder Microchiroptera Dobson, 1875 late early Pliocene in age. 292 S.J. Hand, B.S. Mackness, C.E. Wilkinson, D.M. Wilkinson

B

Figure 1 Mormopterus (Micronomus) sp., from the Pliocene Chinchilla Local Fauna, southeastern Queensland. QM F30575, left upper canine. A, posterior view; B, lingual view; C-C', stereopair, antero-occlusal view. Scale bar = 1 mm.

Description such that the anterolingual blade is convex and the QM F30575 is massively-rooted with a tall, posterolingual crest concave. There is no slender, dominant main cusp (Figure 1). There is a development of a secondary cusp on either blade. narrow cingulum around the whole tooth and a The crown is flattened on the lingual side, and the small but distinct posterolingual cingular cusp. lingual margin is very straight (i.e., in occlusal view There are no other cingular cusps. The basal outline it is neither concave nor convex). The anterobuccal of the crown is subtriangular. The antero- and face is marked by a broad groove which is posterolingual corners of the main cusp (paracone) attenuated towards the apex of the cusp. This are defined by vertical crests; the posterobuccal groove results in a distinct concavity in the surface is convex. The shaft is recurved posteriorly anterobuccal margin of the crown and contributes Pliocene rnolossid from Chinchilla 293 to the tooth's triangular (rather than semi-circular) anterior cingular cusp. It differs from nycteridids in occlusal outline. In occlusal view, there is also a its flattened (rather than concave) lingual face and broad and marked indentation in the posterobuccal lack of anterior cingular cusp, and from margin, perhaps for the following tooth (but see rhinolophids in its complete and conspicuous Remarks below). The massive root is oval in cross­ cingulum. section with a deep groove longitudinally bisecting The Chinchilla canine differs from those of most its flattened lingual face. noctilionoids (phyllostomids, mormoopids and noctilionids) in having a distinctive anterior groove. Remarks However, some phyllostomines (e.g., P!zyllostomus and Trac!zops species) do possess a broad groove on Comparisons Made the anterior face of the shaft as well as a less Its small size, complete cingulum, flattened developed one on the anterobuccal face. But these lingual face and lack of posterior heel suggest that are large species with much larger canines, which QM F30575 is the upper canine of a usually also differ in having a rather bilobed lingual microchiropteran bat. Megachiropteran canines are margin. Many stenodermatines differ in having a generally larger, lack a complete cingulum (absent dorsal expansion of the buccal face of the canine. In buccally, variably developed lingually and Vampyrum the shaft is flattened in front but, like the posteriorly) and are generally not subtriangular in Chinchilla tooth, it has a moderately developed cross-section. Many megachiropteran canines (e.g., cingulum and distinct posterolingual cingular cusp. Pteropus spp.) have very conspicuous ridges on the In most phyllostomines (e.g., Micronycteris), the shaft; in others a secondary cusp is formed by slight upper canine is simple with the anterior shaft elongation of the cingulum (e.g., Nyctimene spp.) or smoothly rounded and with a well-developed is developed on the posterior edge of the shaft (e.g., cingulum but no extra cusps. In desmodontine species of Pteralopex and Harpyionycteris; the latter phyllostomids, the canine is large, long and narrow, may have well-developed posterior and lingual with an acute point and very sharp cutting edge. cingula and/or posterolingual basal cusps). Like the In the other phyllostomid subfamilies (America's Chinchilla canine, species of Macroglossus, glossophagines, stenodermatines, phyllonycterines) Syconycteris, Notopteris and Eonycteris have a deep there is even more variation in the basic form of the longitudinal groove on the anterior face of the shaft upper canine, making generalizations difficult. (in Melonycteris the groove is bounded by sharp Miller (1907) has described the dentition of many ridges), but otherwise differ strikingly from the taxa in this speciose group, and the following fossil in their very weak and incomplete cingula comparisons with the Chinchilla canine can be and rounded or square basal outline. made. In glossophagines, the canine tends to be The Chinchilla canine has been compared with simple with a slightly developed cingulum and no representatives of all microchiropteran groups, secondary cusps, though in some species (e.g., except the monotypic Craseonycteridae and Leptonycteris) the cingulum is obsolete but Tomopeatinae. It differs strikingly from represented anteriorly and posteriorly by short, emballonurid and rhinopomatid canines in its well-developed cusps. In other species (e.g., subtriangular occlusal outline, anterior groove and Sturnira) the inner surface is strongly concave and lack of well-developed anterior and posterior cusps thus unlike the Chinchilla tooth. Stenodermatine and/or cingular cusps. In most emballonuroids canines appear to be quite unlike the Chinchilla (e.g., R!zinopoma, Emballonura and New World tooth. In many stenodermatines, the canine is nearly diclidurines), there is no distinct cingulum, lateral as long as tall, the cingulum is well developed and compression is marked, heel development common, there are no secondary cusps; in some species (e.g., and anterior and posterior cusps are developed at Ardops, P!zyllops and Ariteus) the canine is only just the base of the anterior and posterior cutting edges. taller than the large premolar. In other The upper canines of species of Taphozous and stenodermatines (e.g., Brac!zyp!zylla) the lingual Saccolaimus have very large heels and deep, well­ cingulum forms two low but very distinct cusps developed lingual cingula, often with pronounced (rarely, a secondary cusp extends nearly to the lingual cusps. In lateral view, emballonuroid middle of the posterior edge). Some taxa (e.g., canines are typically more recurved posteriorly than Pygoderma) have a wide posterolingual heel and the Chinchilla canine. minute posterior cingular cusp, others (e.g., The fossil differs from rhinolophoid (rhinolophid, Centurio) are flattened antero-posteriorly (rather hipposiderid, megadermatid and nycteridid) upper than laterally). In phyllonycterines, the canine is canines in its subtriangular (rather than laterally simple but has a poorly defined cingulum, without compressed) crown and its anterior groove. secondary cusps and with the lingual face slightly Additionally, it differs from megadermatids and concave. many hipposiderids in lacking a posterior In noctilionids the canine is short (antero­ secondary cusp at the base of the shaft and an posteriorly) and very tall with a distinctly oblique 294 S.J. Hand, B.S. Mackness, C.E. Wilkinson, D.M. Wilkinson cingulum without secondary cusps, the shaft's complete cingulum and is of similar proportions to lingual surface slightly concave with a median the Chinchilla canine. There is no posterobuccal ridge, the outer surface uniformly convex. In indentation in basal outline. mormoopids, the canine has a conspicuously Only molossid canines characteristically exhibit a concave lingual surface, distinct anterior and broad and distinct groove on the anterior surface of posterior cutting edges and a fairly well developed the shaft, as in the Chinchilla canine. This groove is cingulum (though weak buccally) without found in both Old and New World molossids secondary cusps. including species of Mormopterus, Micronomus, The Chinchilla canine more closely resembles the Sauromys, Platymops, Chaerephon, Molossops, upper canines of vespertilionoids s.1., mystacinids Cheiromeles and Eumops. It is present but less distinct and molossids. VespertiIionoids s.1. (vespertilionids, in many species of Nyctinomus and Eomops. Most natalids, thyropterids, furipterids, myzopodids) and molossids also share the distinct but small cingulum mystacinids, like most bats, lack the anterior groove and distinctly flattened lingual face seen in the distinguishing the Chinchilla canine. Among Chinchilla tooth. Species of Nyctinomus, Chaerephon vespertilionine canines, three basic types were and Molossops lack noticeable cingular cusps, but a recognized by Menu (1985), none of which matches posterobasal cusp is found in species of Cheiromeles, the morphology of the Chinchilla tooth. In Type A Platymops, Mormoptents, Sauromys and Micronomus. (as in, e.g., Myotis and Lasionycteris species) and Compared with the Chinchilla tooth, Cheiromeles Type C canines (e.g., species of Eptesicus, la, canines are much more robust, while those of Scotophilus, Scoteanax, Otonycteris and Histiotis) the Platymops are short and weak (shaft diameter base of the crown is oval rather than subtriangular. conspicuously less than crown diameter). In Further, Type A canines are simple with a distinct Sauromys the basal outline is lingually concave and though rather small cingulum and no secondary more posterolingually extended, and the posterior cusps; Type C canines are more complex with a cutting edge more pronounced. Species of distinct lingual crest. Histiotus has an anterior Mormoptents (Mormoptents) (e.g., M. minutus, M. groove, but unlike the Chinchilla fossil it is acetabulosus) have lost p2 (or it is reduced to a developed between a strong anterolingual crest and spicule) and, possibly as a consequence, lack the an anterior ridge, and there is a lingual cingular deep posterobuccal indentation in occlusal outline emargination. Type B canines (e.g., Pipistrellus seen in the Chinchilla canine. However, among species) have a subtriangular base more like the Mormoptents species an indentation in posterobasal Chinchilla canine but no anterior groove nor outline does not always accommodate the following posterobasal cusp, the posterior cutting edge often premolar. For example, in Sauromys petrophilus there has an incipient secondary cusp, and an indentation is a distinct posterobuccal indentation but p2 occurs in basal outline occurs posterolingually rather than (more lingually) in the toothrow. Unlike the posterobuccally. The canines of miniopterines (very Chinchilla tooth, the canines of Chaerephon and round, tall and simple), nyctophilines (concave Mops species typically have a marked median lingual face, convex lingual shelf), kerivoulines longitudinal crest bordered by grooves on the (laterally very compressed) and most murinines are lingual face of the shaft. simple in form with distinct but low cingula In overall size, relative length and height, occlusal without cusps, generally with a circular or oval basal outline, extent and postition of groove, degree basal outline. of posterior curvature, development of cingulum, In natalids and myzopodids, like kerivoulines, the size and position of posterocingular cusp, and canine is simple, with a distinct but small cingulum groove in the lingual face of the root, the Chinchilla without secondary cusps, the shaft appearing canine closely resembles the canines of the six or so compressed due to its concave lingual face. living species of Mormopterus (Micronomus) Thyropterids are similar, although lingual cingular distributed across Meganesia. It is less similar to cusps are developed in some species but not others Australia's much larger Nyctinomus australis which (Czaplewski 1997). In some natalids (e.g., species of has a semicircular occlusal outline, a distinct lingual Nyctiellus) and in furipterids, the canine is greatly crest and shelf, and no posterobasal cusp nor reduced in size, being only just taller than the marked indentation for the following tooth. It posterior premolar. Furipterids further differ from differs from the medium-sized Chaerephon jobensis the Chinchilla tooth in having a well-developed in its subtriangular rather than semicircular occlusal secondary cusp on the posterior cutting edge and a outline and in lacking a median crest and grooves well-developed cingulum forming a conspicuous on the lingual face. Canines of Riversleigh's anterior basal cusp and smaller posterior one. Miocene molossids Petramops creaseri Hand, 1990, The upper canine of mystacinids has no distinct and Mormoptents (Hydromops) riversleighensis Hand (but perhaps an incipient) groove, a semicircular et al., 1997, are not yet known, but other species of rather than subtriangular basal outline, and no Hydromops (Revilliod 1920; Legendre 1982; obvious posterobasal cusp. It does, however, have a Legendre et al. 1988) appear to differ from the Pliocene molossid from Chinchilla 295

Chinchilla tooth in characteristically exhibiting a creaseri (Hand 1990) and MOrrllOpterlls (Hydromops) strong medial rib on the buccal face of the crown riversleighensis (Hand et al. 1997), both of which (Engesser 1972; Legendre et al. 1988). appear to lie outside the radiation of living Differences in occlusal basal outline, Australian molossids. They lack several derived development of lingual shelf, and size and position traits exhibited by species of Micronomlls, of posterocingular cusp and posterobasal shelf Nyctinol1llls and Chaerephon, and seem to have been seem to distinguish the various Micronomlls part of an older, archaic bat fauna that was species. In such features, the Chinchilla tooth is widespread, had close relatives in Europe and less similar to M. beccarii, the species found in the North America, and left no living descendants in Chinchilla area today (McKenzie 1995; but see Australia (Hand 1990). Allison 1996), than to other species including M. Legendre (1984a, 1984b, 1984c, 1985) speculated norfolcenis of eastern New South Wales. However, that the genus Mormopterlls was probably derived because intraspecific variation (including sexual at the end of the Eocene or beginning of the dimorphism) may be considerable in molossid bat Oligocene from European species of Cuvieril1lops, canines (e.g., Engesser 1972), it has not been and from there expanded worldwide to wherever possible to refer the Chinchilla canine to any climatic conditions were favourable. Species of particular species of Micronomlls. Morl1lopterlls reached South America by the late Oligocene (i.e., M. (Neol1lops) faustoi; Paula Couto 1956; Paula Couto and Mezzalira 1971), and DISCUSSION Australia by the late Oligocene (i.e., MOrrllOpterus Chinchilla is the seventh Australian locality to (Hydromops) riversleighensis). The latter subgenus, produce Tertiary-aged bats, the other six localities Hydrol1lops, is known also from the early Miocene of being: Murgon, southeastern Queensland (early Thailand (Legendre et al. 1988) and the early to Eocene Tingamarra Local Fauna); Lake middle Miocene of Europe (Revilliod 1920), Palankarinna, central Australia (late Oligocene disappearing from Europe after that time. Close Di~imanka Local Fauna); Riversleigh, northwestern relatives (though not descendants) of the M. Queensland (late Oligocene to Pliocene faunas); (Hydrol1lops) lineage include the living African Bullock Creek, Northern Territory (middle Miocene species M. (Platyl1lops) setiger and Australia's M. Bullock Creek Local Fauna); Hamilton, Victoria (Micronomus) species (Legendre 1984a, 1984b; (early Pliocene Hamilton Local Fauna); and Legendre et al. 1988; Hand et al. 1997). Wellington Caves, central New South Wales (early­ The ancestors of the Australian endemic middle Pliocene Big Sink Local Fauna); (Turnbull Micronol1llls lineage are thought to have arrived in and Lundelius 1970; Archer 1978; Archer et al. 1994; the Australian region from Asia by at least the late Hand et al. 1988, 1994, 1998). From the Australian Oligocene, independently of Petramops, Nyctinomus Pliocene sites, the bat families Hipposideridae, and Chaerephon species (Legendre 1984a, 1984b, Megadermatidae, Emballonuridae and 1984c, 1985; Hand 1990). The Chinchilla molossid Vespertilionidae have been recorded (Turnbull and testifies that the Micronol1lus lineage itself had Lundelius 1970; Hand et al. 1988; Archer et al. 1994; evolved by the middle Pliocene. Today, the four Hand 1995, 1996; Hand and Godthelp this volume). named living Micronomus species (beccarii, loriae, The Chinchilla bat is the first Australian Pliocene norfolcensis and planiceps) and at least two others representative of the cosmopolitan bat family (separable on fur and penis characters; Richards Molossidae. Worldwide, some 80 living molossid 1995) are distributed across the Australian species are referred to between 9 and 12 genera mainland, with two, loriae and beccarii, found also in (Freeman 1981; Honacki et al. 1982; Legendre 1984b) New Guinea, and beccarii occurring on Ambon with another 20 or so fossil (Tertiary) taxa reported island (McKenzie 1995). from all continents except Antarctica (Legendre Micronol1lus species, like other molossids, are fast, 1985; Hand 1990; Czaplewski 1997). In Australia direct fliers which forage for flying insects in open today, the Molossidae is represented by one species areas above the canopy or over open water and each of the genera Nyctinomlls and Chaerephon and typically roost in tree hollows, rock crevices and at least six species of Mormopterlls (Micronol1l!ls) caves. In Australia, they occur in desert, semi-arid whose taxonomy remains confused (Allison 1989, and mesic regions, foraging over rainforests, 1996; Parnaby 1991; Richards 1995). mangroves, open woodland and savanna (Strahan Currently, Australian Tertiary molossids are 1995). All Riversleigh molossid specimens have known only from late Oligocene and early Miocene been recovered from karstic sediments which deposits at Riversleigh in northwestern appear to have accumulated within closed forest Queensland, and now from Chinchilla. They are not (Hand 1990; Hand et al. 1997). However, molossids yet known from Riversleigh's Pliocene Rackham's are rare at Riversleigh compared with other bat taxa Roost Site (Archer et al. 1994). The Riversleigh (especially rhinolophoids) and it is not yet clear molossids represent at least two species, Petramops whether they lived in caves or forest, or were 296 S.]. Hand, B.S. Mackness, CE. Wilkinson, D.M. Wilkinson brought in as prey from adjacent habitats by Archer, M. and Bartholomai, A. (1978). Tertiary carnivorous megadermatids (Hand 1990). In mammals of Australia: a synoptic review. Alcheringa Europe, Tertiary molossids have been recorded as 2: 1-19. whole skeletons from strictly lacustrine deposits Archer, M and Dawson, L. (1982). Revision of marsupial (Sige 1971). lions of the genus Thylacoleo Gervais (Thylacoleon­ While the exact nature of the Chinchilla idae, Marsupialia) and thylacoleonid evolution in the palaeoenvironment has yet to be interpreted, the late Cainozoic. In Archer, M. (ed.), Carnivorous nature of the sediments suggests a fluviatile rather marsupials, 2: 477-494, Surrey Beatty & Sons, Chipping Norton, N.5.W. than lacustrine depositional environment. The composition of the mammalian fauna suggests a Archer, M., Hand, S.J. and Godthelp, H. (1994). Riversleigh: the story of animals in Australia's ancient woodland environment with a lack of any inland rainforests, 1-264, reprint with revisions, Reed undoubted closed-forest taxa. The predominance of Books, Chatswood, N.5.W. grazing kangaroos and diprotodontids indicates Bartholomai, A (1962). A new species of Thylacoleo and that grasslands were part of the environment. The notes on some caudal vertebrae of Palorchestes azael. presence of a molossid bat does not contradict that Memoirs ofthe Queensland Museum 14: 33-40. interpretation. Bartholomai, A. (1963). Revision of the extinct macropodid genus Sthenurus Owen in Queensland. Memoirs ofthe Queensland Museum 14: 51-76. ACKNOWLEDGEMENTS Bartholomai, A (1966). The type specimens of some of de The study of the Chinchilla material was Vis' species of fossil Macropodidae. Memoirs of the supported in part by an ARC Program Grant to M. Queensland Museum 14: 115-126. Archer; a grant from the Department of Arts, Sport, Bartholomai, A (1967). Troposodon, a new genus of fossil the Environment, Tourism and Territories to M. Macropodinae (Marsupialia). Memoirs of the Archer, S.J. Hand and H. Godthelp; a grant from Queensland Museum 15: 21-33. the National Estate Program Grants Scheme to M. Bartholomai, A (1971). DasyuTtls dunmalli, a new species Archer and A. Bartholomai; and grants in aid to the of fossil marsupial (Dasyuridae) in the upper Riversleigh Research Project from the University of Cainozoic deposits of Queensland. Memoirs of the New South Wales, Wang Australia Pty Ltd, ICI Queensland Museum 16: 19-26. Australia and the Australian Geographic Society. Bartholomai, A (1973). 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